For successful locomotion, animals require visual and somatosensory information. Even though a number of supraspinal centers receive both in varying degrees, processing this information at different levels of the central nervous system, especially their contribution to visuo-motor and sensory-motor integration during locomotion is poorly understood.
This dissertation investigates the patterns of neuronal activity in three areas of the forebrain in the cat performing different locomotor tasks to elucidate involvement of these areas in processing of visual and somatosensory information related to locomotion. In three studies, animals performed two contrasting locomotor tasks in each and the neuronal activities were analyzed.
In the first study, cats walked in either complete darkness or in an illuminated room while the neuronal activity of the motor cortex was recorded. This study revealed that the neuronal discharge patterns in the motor cortex were significantly different between the two illumination conditions. The mean discharge rates, modulation, and other variables were significantly different in 49% of the neurons. This suggests a contextual correlation between the motor cortical activity and being able to see.
In two other studies, the activities of neurons of either the somatosensory cortex (SI) or ventrolateral thalamus (VL) were recorded while cats walked on a flat surface (simple locomotion) or along a horizontal ladder where continuous visual and somatosensory feedback was required (complex locomotion).
We found that the activity of all but one SI cells with receptive fields on the sole peaked before the foot touched the ground: predictably. Other cells showed various patterns of modulation, which differed between simple and complex locomotion. We discuss the predictive and reflective functionality of the SI in cyclical sensory-motor events such as locomotion.
We found that neuronal discharges in the VL were modulated to the stride cycle resembling patterns observed in the cortex that receives direct inputs from the VL. The modulation was stronger during walking on the ladder revealing VL’s contribution to locomotion-related activity of the cortex during precision stepping.
Distinct neuronal adaptations between simple and complex locomotion tasks were observed for neurons with different receptive field properties and fast- and slow-conducting pyramidal tract neurons. Feedforward and feedback-driven kinematic control strategies were observed for adaptations to expected and unexpected perturbations, respectively, during complex locomotion tasks. These kinematic differences were reflected in the response characteristics of motor cortical neurons receptive to somatosensory information from different parts of the forelimb, elucidating roles for the various neuronal populations in accommodating disturbances in the environment during behaviors. The results show that anatomical and physiological characteristics of motor cortical neurons are important for determining if and how neurons are involved in precise control of locomotion during natural behaviors.
Background: Directional preferences during center-out horizontal shoulder-elbow movements were previously established for both the dominant and non-dominant arm with the use of a free-stroke drawing task that required random selection of movement directions. While the preferred directions were mirror-symmetrical in both arms, they were attributed to a tendency specific for the dominant arm to simplify control of interaction torque by actively accelerating one joint and producing largely passive motion at the other joint. No conclusive evidence has been obtained in support of muscle effort minimization as a contributing factor to the directional preferences. Here, we tested whether distal load changes directional preferences, making the influence of muscle effort minimization on the selection of movement direction more apparent.
Methods: The free-stroke drawing task was performed by the dominant and non-dominant arm with no load and with 0.454 kg load at the wrist. Motion of each arm was limited to rotation of the shoulder and elbow in the horizontal plane. Directional histograms of strokes produced by the fingertip were calculated to assess directional preferences in each arm and load condition. Possible causes for directional preferences were further investigated by studying optimization across directions of a number of cost functions.
Results: Preferences in both arms to move in the diagonal directions were revealed. The previously suggested tendency to actively accelerate one joint and produce passive motion at the other joint was supported in both arms and load conditions. However, the load increased the tendency to produce strokes in the transverse diagonal directions (perpendicular to the forearm orientation) in both arms. Increases in required muscle effort caused by the load suggested that the higher frequency of movements in the transverse directions represented increased influence of muscle effort minimization on the selection of movement direction. This interpretation was supported by cost function optimization results.
Conclusions: While without load, the contribution of muscle effort minimization was minor, and therefore, not apparent, the load revealed this contribution by enhancing it. Unlike control of interaction torque, the revealed tendency to minimize muscle effort was independent of arm dominance.